Bumblebee Family PDF

8/15/2020 Bumblebee - Wikipedia
Bumblebee
A bumblebee (or bumble bee, bumble-bee, or humble-bee)
is any of over 250 species in the genus Bombus, part of Apidae, Bumblebee
one of the bee families. This genus is the only extant group in the Temporal range: Eocene–Present
tribe Bombini, though a few extinct related genera (e.g., PreꞒ Ꞓ OS D C P T J K PgN
Calyptapis) are known from fossils. They are found primarily in
higher altitudes or latitudes in the Northern Hemisphere, although
they are also found in South America where a few lowland tropical
species have been identified. European bumblebees have also been
introduced to New Zealand and Tasmania. Female bumblebees can
sting repeatedly, but generally ignore humans and other animals.
Most bumblebees are social insects that form colonies with a single
queen. The colonies are smaller than those of honey bees, growing
to as few as 50 individuals in a nest. Cuckoo bumblebees are brood
parasitic and do not make nests; their queens aggressively invade Buff-tailed bumblebee (Bombus
the nests of other bumblebee species, kill the resident queens and terrestris)
then lay their own eggs, which are cared for by the resident
workers. Cuckoo bumblebees were previously classified as a Scientific classification
separate genus, but are now usually treated as members of Kingdom: Animalia
Bombus.
Phylum: Arthropoda
Bumblebees have round bodies covered in soft hair (long branched Class: Insecta
setae) called pile, making them appear and feel fuzzy. They have
aposematic (warning) coloration, often consisting of contrasting Order: Hymenoptera
bands of colour, and different species of bumblebee in a region
Family: Apidae
often resemble each other in mutually protective Müllerian
mimicry. Harmless insects such as hoverflies often derive Tribe: Bombini
protection from resembling bumblebees, in Batesian mimicry, and
Genus: Bombus
may be confused with them. Nest-making bumblebees can be
distinguished from similarly large, fuzzy cuckoo bees by the form Latreille, 1802
of the female hind leg. In nesting bumblebees, it is modified to Diversity
form a pollen basket, a bare shiny area surrounded by a fringe of
hairs used to transport pollen, whereas in cuckoo bees, the hind leg > 250 species and subspecies
is hairy all round, and they never carry pollen.
Like their relatives the honeybees, bumblebees feed on nectar,

using their long hairy tongues to lap up the liquid; the proboscis is
folded under the head during flight. Bumblebees gather nectar to
add to the stores in the nest, and pollen to feed their young. They
forage using colour and spatial relationships to identify flowers to
feed from. Some bumblebees steal nectar, making a hole near the Natural distribution shown in red.
base of a flower to access the nectar while avoiding pollen transfer. Introductions to New Zealand, spread
Bumblebees are important agricultural pollinators, so their decline to Tasmania not shown
in Europe, North America, and Asia is a cause for concern. The
decline has been caused by habitat loss, the mechanisation of agriculture, and pesticides.
https://en.wikipedia.org/wiki/Bumblebee 1/28
Contents
Etymology and common names
Phylogeny
Taxonomy
General description
Distribution and habitat
Biology
Feeding
Wax production
Coloration
Temperature control
Chill-coma temperature
Communication and social learning
Reproduction and nesting
Foraging behaviour
Asynchronous flight muscles
Cuckoo bumblebees
Sting
Predators, parasites, and pathogens
Relationship to humans
Agricultural use
Population decline
Conservation efforts
Misconception about flight
In music and literature
Military
See also
Notes
References
Sources
External links
Etymology and common names

The word "bumblebee" is a compound of "bumble" + "bee"—"bumble" meaning to hum, buzz, drone, or
move ineptly or flounderingly.[1] The generic name Bombus, assigned by Pierre André Latreille in 1802,
is derived from the Latin word for a buzzing or humming sound, borrowed from Ancient Greek βόμβος
(bómbos).[2][3]
According to the Oxford English Dictionary (OED), the term "bumblebee"

was first recorded as having been used in the English language in the 1530
work Lesclarcissement by John Palsgrave, "I bomme, as a bombyll bee
dothe."[4] However the OED also states that the term "humblebee"
predates it, having first been used in 1450 in Fysshynge wyth Angle, "In
Juyll the greshop & the humbylbee in the medow."[5] The latter term was
used in A Midsummer Night's Dream (circa 1600) by William Shakespeare,
"The honie-bags steale from the humble Bees."[6] Similar terms are used in
other Germanic languages, e.g. German Hummel (OHG humbala[7]), Dutch
hommel or Swedish humla.
An old provincial name, "dumbledor", also denoted a buzzing insect such as

a bumblebee or cockchafer, "dumble" probably imitating the sound of these Beatrix Potter called
Babbity Bumble a
insects, while "dor" meant "beetle".[8]
"bumble bee" in The Tale of
Mrs Tittlemouse, 1910.
In On the Origin of Species (1859), Charles Darwin speculated about
"humble-bees" and their interactions with other species:[9]
I have [...] reason to believe that humble-bees are indispensable to the fertilisation of the
heartsease (Viola tricolor), for other bees do not visit this flower. From experiments which I
have tried, I have found that the visits of bees, if not indispensable, are at least highly
beneficial to the fertilisation of our clovers; but humble-bees alone visit the common red
clover (Trifolium pratense), as other bees cannot reach the nectar.
However, "bumblebee" remained in use, for example in The Tale of Mrs. Tittlemouse (1910) by Beatrix
Potter, "Suddenly round a corner, she met Babbitty Bumble--"Zizz, Bizz, Bizzz!" said the bumble bee."
Since World War II "humblebee" has fallen into near-total disuse.[10]
Phylogeny
The bumblebee tribe Bombini is one of four groups of corbiculate bees (those with pollen baskets) in the
Apidae, the others being the Apini (honey bees), Euglossini (orchid bees), and Meliponini (stingless
bees). The corbiculate bees are a monophyletic group. Advanced eusocial behaviour appears to have
evolved twice in the group, giving rise to controversy, now largely settled, as to the phylogenetic origins
of the four tribes; it had been supposed that eusocial behaviour had evolved only once, requiring the
Apini to be close to the Meliponini, which they do not resemble. It is now thought that the Apini (with
advanced societies) and Euglossini are closely related, while the primitively eusocial Bombini are close to
the Meliponini, which have somewhat more advanced eusocial behaviour. Sophie Cardinal and Bryan
Danforth comment that "While remarkable, a hypothesis of dual origins of advanced eusociality is
congruent with early studies on corbiculate morphology and social behavior."[11] Their analysis,
combining molecular, morphological and behavioural data, gives the following cladogram:[11]
Corbiculate bees
 
  Apini (honeybees)
 
 
 
  Euglossini (orchid bees)
 
   
  Bombini (bumblebees)
   
Meliponini (stingless bees)
 
On this hypothesis, the molecular data suggest that the Bombini are 25 to 40 million years old, while the
Meliponini (and thus the clade that includes the Bombini and Meliponini) are 81 to 96 million years old,
about the same age as the corbiculate group.[11]
However, a more recent phylogeny using transcriptome data from 3,647 genes of ten corbiculate bee
species supports the single origin of eusociality hypothesis in the corbiculate bees.[12] They find that
Bombini is in fact sister to Meliponini, corroborating that previous finding from Sophie Cardinal and
Bryan Danforth (2011). However, Romiguier et al. (2015) shows that Bombini, Meliponini, and Apini
form a monophyletic group, where Apini shares a most recent common ancestor with the Bombini and
Meliponini clade, while Euglossini is most distantly related to all three, since it does not share the same
most recent common ancestor as Bombini, Meliponini, and Apini. Thus, their analysis supports the
single origin of eusociality hypothesis within the corbiculate bees, where eusociality evolved in the
common ancestor of Bombini, Apini, and Meliponini.
The fossil record for bees is incomplete. Around 11 specimens that might possibly be Bombini, some
poorly documented, had been described by 2011; some (such as Calyptapis florissantensis from
Florissant, USA, and Oligoapis beskonakensis from Beskonak, Turkey) dated from the Oligocene.[13] In
2012 a fossil bumblebee, Bombus (Bombus) randeckensis was described from the Miocene Randeck
Maar in southwestern Germany and confidently placed in the subgenus Bombus.[13] In 2014, another
species, Bombus cerdanyensis, was described from Late Miocene lacustrine beds of La Cerdanya, Spain,
but not placed into any subgenus,[14] while a new genus and species, Oligobombus cuspidatus was
described from the late Eocene Bembridge Marls of the Isle of Wight.[15][16] The species Bombus
trophonius was described in October 2017 and placed in Bombus subgenus Cullumanobombus.[17]
Taxonomy
The genus Bombus, the only one extant genus in the tribe Bombini, comprises over 250 species;[18] for
an overview of the differences between bumblebees and other bees and wasps, see characteristics of
common wasps and bees. The genus has been divided variously into up to 49 subgenera, a degree of
complexity criticised by Williams (2008).[19] The cuckoo bumblebees Psithyrus have sometimes been
treated as a separate genus but are now considered to be part of Bombus, in one or more subgenera.[19]
Examples of Bombus species include Bombus atratus, Bombus dahlbomii, Bombus fervidus, Bombus
lapidarius, Bombus ruderatus, and Bombus rupestris.
Bombus (genus)  
      Mendacibombus, 12 species
 
 
 
Bombias, 3 species
 
    Kallobombus, 1 species
 
 
    Orientalibombus, 3 species
 
 
  Subterraneobombus, 10 species
 
 
Megabombus, 22 species
   
 
Thoracobombus, 50 species
 
 
 
 
 
Psithyrus, 30 species
 
 
Pyrobombus, 50 species
   
 
Alpinobombus, 5 species
 
 
 
 
 
Bombus (subgenus), 5 species
 
 
 
Alpigenobombus, 7 species
 
 
 
Melanobombus, 17 species
 
 
 
 
Sibiricobombus, 7 species
 
 
 
 
Cullumanobombus, 23 species
Subgenera of the genus Bombus
General description
Bumblebees vary in appearance, but are generally plump and densely furry. They are larger, broader and
stouter-bodied than honeybees, and their abdomen tip is more rounded. Many species have broad bands
of colour, the patterns helping to distinguish different species. Whereas honeybees have short tongues
and therefore mainly pollinate open flowers, some bumblebee species have long tongues and collect
nectar from flowers that are closed into a tube.[20] Bumblebees have fewer stripes (or none), and usually
have part of the body covered in black fur, while honeybees have many stripes including several grey
stripes on the abdomen.[21] Sizes are very variable even within species; the largest British species, B.
terrestris, has queens up to 22 mm (0.9 in) long, males up to 16 mm (0.6 in) long, and workers between
11 and 17 mm (0.4–0.7 in) long.[22] The largest bumblebee species in the world is B. dahlbomii of Chile,
up to about 40 mm (1.6 in) long, and described as "flying mice" and "a monstrous fluffy ginger beast".[23]
Distribution and habitat

Bumblebees are typically found in temperate climates, and are often found at higher latitudes and
altitudes than other bees, although a few lowland tropical species exist.[24] A few species (B. polaris and
B. alpinus) range into very cold climates where other bees might not be found; B. polaris occurs in
northern Ellesmere Island in the high Arctic, along with another bumblebee B. hyperboreus, which
parasitises its nest. This is the northernmost occurrence of any eusocial insect.[25] One reason for their
presence in cold places is that bumblebees can regulate their body temperature, via solar radiation,
internal mechanisms of "shivering" and radiative cooling from the abdomen (called heterothermy).
Other bees have similar physiology, but the mechanisms seem best developed and have been most
studied in bumblebees.[26] They adapt to higher elevations by extending their wing stroke amplitude.[27]
Bumblebees have a largely cosmopolitan distribution but are absent from Australia (apart from
Tasmania where they have been introduced) and are found in Africa only north of the Sahara.[28] More
than a hundred years ago they were also introduced to New Zealand, where they play an important role
as efficient pollinators.
Biology
Feeding
The bumblebee tongue (the proboscis) is a long, hairy structure that extends
from a sheath-like modified maxilla. The primary action of the tongue is
lapping, that is, repeated dipping of the tongue into liquid.[29] The tip of the
tongue probably acts as a suction cup and during lapping, nectar may be
drawn up the proboscis by capillary action. When at rest or flying, the
proboscis is kept folded under the head. The longer the tongue, the deeper
the bumblebee can probe into a flower and bees probably learn from A common carder
experience which flower source is best-suited to their tongue length.[30] bumblebee Bombus
Bees with shorter proboscides, like Bombus bifarius, have a more difficult pascuorum extending its
time foraging nectar relative to other bumblebees with longer proboscides; tongue towards a Heuchera
to overcome this disadvantage, B. bifarius workers were observed to lick the inflorescence
back of spurs on the nectar duct, which resulted in a small reward.[31]
Wax production
The exoskeleton of the abdomen is divided into plates called dorsal tergites and ventral sternites. Wax is
secreted from glands on the abdomen and extruded between the sternites where it resembles flakes of
dandruff. It is secreted by the queen when she starts a nest and by young workers. It is scraped from the
abdomen by the legs, moulded until malleable and used in the construction of honeypots, to cover the
eggs, to line empty cocoons for use as storage containers and sometimes to cover the exterior of the
nest.[32]
Coloration
The brightly coloured pile of the bumblebee is an aposematic (warning) signal, given that females can
inflict a painful sting. Depending on the species and morph, the warning colours range from entirely
black, to bright yellow, red, orange, white, and pink.[33] Dipteran flies in the families Syrphidae
(hoverflies), Asilidae (robber flies), Tabanidae (horseflies), Oestridae (bot or warble flies) and
Bombyliidae (bee flies, such as Bombylius major) all include Batesian mimics of bumblebees, resembling
them closely enough to deceive at least some predators.[34]
Many species of Bombus, including the group sometimes called Psithyrus

(cuckoo bumblebees), have evolved Müllerian mimicry, where the different
bumblebees in a region resemble each other, so that a young predator need
only learn to avoid any of them once. For example, in California a group of
bumblebees consists of largely black species including B. californicus, B.
caliginosus, B. vandykei, B. vosnesenskii, B. insularis and B. fernaldae.
Other bees in California include a group of species all banded black and
yellow. In each case, Müllerian mimicry provides the bees in the group with
a selective advantage.[34] In addition, parasitic (cuckoo) bumblebees
resemble their hosts more closely than would be expected by chance, at
least in areas like Europe where parasite-host co-speciation is common; but
this too may be explained as Müllerian mimicry, rather than requiring the
parasite's coloration to deceive the host (aggressive mimicry).[35]
Cuckoo bumblebees, like

Temperature control this Bombus barbutellus,
have similar aposematic
Bumblebees are active under conditions during which honeybees stay at (warning) coloration to nest-
home, and can readily absorb heat from even weak sunshine.[36] The thick making bumblebees, and
pile created by long setae (bristles) acts as insulation to keep bumblebees may also mimic their host
warm in cold weather; species from cold climates have longer setae (and species.
thus thicker insulation) than those from the tropics.[37] The temperature of
the flight muscles, which occupy much of the thorax, needs to be at least
30 °C (86 °F) before flight can take place. The muscle temperature can be raised by shivering. It takes
about five minutes for the muscles to reach this temperature at an air temperature of 13 °C (55 °F).[38]
Chill-coma temperature
The chill-coma temperature in relation to flying insects is the temperature at which flight muscles cannot
be activated. Compared to honey bees and carpenter bees, bumblebees have the lowest chill-coma
temperature. Of the bumblebees Bombus bimaculatus has the lowest at 7 °C (45 °F). However,
bumblebees have been seen to fly in colder ambient temperatures. This discrepancy is likely because the
chill-coma temperature was determined by tests done in a laboratory setting. However, bumblebees live
in insulated shelters and can shiver to warm up before venturing into the cold.[39]
Communication and social learning
Bumblebees do not have ears, and it is not known whether or how well they can hear. However, they are
sensitive to the vibrations made by sound travelling through wood or other materials.[32]
Bumblebees do not exhibit the "bee dances" used by honeybees to tell other workers the locations of food
sources. Instead, when they return from a successful foraging expedition, they run excitedly around in
the nest for several minutes before going out to forage once more. These bees may be offering some form
of communication based on the buzzing sounds made by their wings, which may stimulate other bees to
start foraging.[40] Another stimulant to foraging activity is the level of food reserves in the colony. Bees
monitor the amount of honey in the honeypots, and when little is left or when high quality food is added,
they are more likely to go out to forage.[41]
Bumblebees have been observed to partake in social learning. In a 2017 study involving Bombus
terrestris, bees were taught to complete an unnatural task of moving large objects to obtain a reward.
Bees that first observed another bee complete the task were significantly more successful in learning the
task than bees that observed the same action performed by a magnet, indicating the importance of social
information. The bees did not copy each other exactly: in fact, the study suggested that the bees were
instead attempting to emulate each other's goals.[42][43]
Reproduction and nesting
Nest size depends on species of bumblebee. Most form colonies of

between 50 and 400 individuals,[44] but colonies have been
documented as small as ~20 individuals and as large as 1700.[45]
These nests are small compared to honeybee hives, which hold about
50,000 bees. Many species nest underground, choosing old rodent
burrows or sheltered places, and avoiding places that receive direct
sunlight that could result in overheating. Other species make nests
above ground, whether in thick grass or in holes in trees. A
bumblebee nest is not organised into hexagonal combs like that of a
Nest of red-tailed bumblebee.
honeybee; the cells are instead clustered together untidily. The
Bombus lapidarius, showing wax
workers remove dead bees or larvae from the nest and deposit them pots full of honey
outside the nest entrance, helping to prevent disease. Nests in
temperate regions last only for a single season and do not survive the
winter.[44]
In the early spring, the queen comes out of diapause and finds a suitable place to create her colony. Then
she builds wax cells in which to lay her eggs which were fertilised the previous year. The eggs that hatch
develop into female workers, and in time, the queen populates the colony, with workers feeding the
young and performing other duties similar to honeybee workers. In temperate zones, young queens
(gynes) leave the nest in the autumn and mate, often more than once, with males (drones) that are
forcibly driven out of the colony.[46] The drones and workers die as the weather turns colder; the young
queens feed intensively to build up stores of fat for the winter. They survive in a resting state (diapause),
generally below ground, until the weather warms up in the spring with the early bumblebee being the
species that is among the first to emerge.[46][47][48] Many species of bumblebee follow this general trend
within the year. Bombus pensylvanicus is a species that follows this type of colony cycle.[49] For this
species the cycle begins in February, reproduction starts in July or August, and ends in the winter
months. The queen remains in hibernation until spring of the following year in order to optimize
conditions to search for a nest.[50]
In fertilised queens, the ovaries only become active when the queen starts to lay. An egg passes along the
oviduct to the vagina where there is a chamber called the spermatheca, in which the sperm from the
mating is stored. Depending on need, she may allow her egg to be fertilised. Unfertilised eggs become
haploid males; fertilised eggs grow into diploid females and queens.[52] The hormones that stimulate the
development of the ovaries are suppressed in female worker bees, while the queen remains dominant.[46]
To develop, the larvae must be fed both nectar for carbohydrates and pollen for protein. Bumblebees feed
nectar to the larvae by chewing a small hole in the brood cell into which they regurgitate nectar. Larvae
are fed pollen in one of two ways, depending on the bumblebee species. Pocket-making bumblebees
create pockets of pollen at the base of the brood-cell clump from which the larvae feed themselves.
Pollen-storing bumblebees keep pollen in separate wax pots and feed it to the larvae.[53]
After the emergence of the

first or second group of
offspring, workers take
over the task of foraging
and the queen spends most
of her time laying eggs and
caring for larvae. The
colony grows progressively
larger and eventually
Bumblebee life-cycle showing adults and begins to produce males
An above-ground nest, hidden in
larvae in nest of B. terrestris. Engraved in and new queens.[46] grass and moss, of the common
1840 by William Home Lizars after drawing Bumblebee workers can lay carder bee, Bombus pascuorum.
probably by James Hope Stewart. [51] unfertilised haploid eggs The wax canopy or involucrum has
(with only a single set of been removed to show winged
chromosomes) that workers and pupae in irregularly
develop into viable male bumblebees. Only fertilised queens can lay placed wax cells.
diploid eggs (one set of chromosomes from a drone, one from the
queen) that mature into workers and new queens.[54]
In a young colony, the queen minimises reproductive competition from workers by suppressing their
egg-laying through physical aggression and pheromones.[55] Worker policing leads to nearly all eggs laid
by workers being eaten.[56] Thus, the queen is usually the mother of all of the first males laid. Workers
eventually begin to lay male eggs later in the season when the queen's ability to suppress their
reproduction diminishes.[57] Because of the reproductive competition between workers and the queen,
bumblebees are considered "primitively eusocial".[11][56]
Although a large majority of bumblebees follow such monogynous colony cycles that only involve one
queen, some select Bombus species (such as Bombus atratus) will spend part of their life cycle in a
polygynous phase (have multiple queens in one nest during these periods of polygyny).[58]
Foraging behaviour
Bumblebees generally visit flowers that exhibit the bee pollination

syndrome and these patches of flowers may be up to 1–2 km from
their colony.[59] They tend to visit the same patches of flowers every
day, as long as they continue to find nectar and pollen there,[60] a
habit known as pollinator or flower constancy. While foraging,
bumblebees can reach ground speeds of up to 15 m/s (54 km/h).[61]
Bumblebees use a combination of colour and spatial relationships to

learn which flowers to forage from.[62] They can also detect both the
A bumblebee loaded with pollen in
presence and the pattern of electric fields on flowers, which occur
its pollen baskets
due to atmospheric electricity, and take a while to leak away into the
ground. They use this information to find out if a flower has been
recently visited by another bee.[63] Bumblebees can detect the
temperature of flowers,[64] as well as which parts of the flower are hotter or cooler[65] and use this
information to recognise flowers. After arriving at a flower, they extract nectar using their long tongues
("glossae") and store it in their crops. Many species of bumblebees also exhibit "nectar robbing": instead
of inserting the mouthparts into the flower in the normal way, these bees bite directly through the base of
the corolla to extract nectar, avoiding pollen transfer.[66]
Pollen is removed from flowers deliberately or incidentally by

bumblebees. Incidental removal occurs when bumblebees come in
contact with the anthers of a flower while collecting nectar. When it
enters a flower, the bumblebee's body hairs receive a dusting of
pollen from the anthers. In queens and workers this is then
groomed into the corbiculae (pollen baskets) on the hind legs
where it can be seen as bulging masses that may contain as many
as a million pollen grains. Male bumblebees do not have corbiculae Biting open the stem of a
and do not purposively collect pollen.[67] Bumblebees are also flower...
capable of buzz pollination, in which they dislodge pollen from the
anthers by creating a resonant vibration with their flight
muscles.[68]
In at least some species, once a bumblebee has visited a flower, it

leaves a scent mark on it. This scent mark deters bumblebees from
visiting that flower until the scent degrades.[69] This scent mark is
a general chemical bouquet that bumblebees leave behind in
different locations (e.g. nest, neutral, and food sites),[70] and they
learn to use this bouquet to identify both rewarding and
unrewarding flowers,[71] and may be able to identify who else has
visited a flower.[72] Bumblebees rely on this chemical bouquet
more when the flower has a high handling time, that is, where it ...and using its tongue to
takes a longer time for the bee to find the nectar once inside the drink the nectar.
flower.[73]
Once they have collected nectar and pollen, female workers return
to the nest and deposit the harvest into brood cells, or into wax
cells for storage. Unlike honeybees, bumblebees only store a few
days' worth of food, so are much more vulnerable to food
shortages.[74] Male bumblebees collect only nectar and do so to A bumblebee "nectar robbing" a flower
feed themselves. They may visit quite different flowers from the
workers because of their different nutritional needs.[75]
Asynchronous flight muscles
Bees beat their wings about 200 times a second. Their thorax muscles do not contract on each nerve
firing, but rather vibrate like a plucked rubber band. This is efficient, since it lets the system consisting of
muscle and wing operate at its resonant frequency, leading to low energy consumption. Further, it is
necessary, since insect motor nerves generally cannot fire 200 times per second.[76] These types of
muscles are called asynchronous muscles[77] and are found in the insect wing systems in families such as
Hymenoptera, Diptera, Coleoptera, and Hemiptera.[76] Bumblebees must warm up their bodies
considerably to get airborne at low ambient temperatures. Bumblebees can reach an internal thoracic
temperature of 30 °C (86 °F) using this method.[26][78]
Cuckoo bumblebees
Bumblebees of the subgenus Psithyrus (known as 'cuckoo bumblebees', and formerly considered a
separate genus) are brood parasites,[79] sometimes called kleptoparasites,[80] in the colonies of other
bumblebees, and have lost the ability to collect pollen. Before finding and invading a host colony, a
Psithyrus female, such as that of the Psithyrus species of B.

sylvestris,[81] feeds directly from flowers. Once she has infiltrated a
host colony, the Psithyrus female kills or subdues the queen of that
colony, and uses pheromones and physical attacks to force the
workers of that colony to feed her and her young.[82] Usually, cuckoo
bumblebees can be described as queen-intolerant inquilines, since
the host queen is often killed to enable the parasite to produce more
offspring,[79] though some species, such as B. bohemicus, actually
enjoy increased success when they leave the host queen alive.[83]
The female Psithyrus has a number of morphological adaptations for

combat, such as larger mandibles, a tough cuticle and a larger venom
The cuckoo bumblebee B. vestalis,
sac that increase her chances of taking over a nest.[84] Upon
a parasite of B. terrestris
emerging from their cocoons, the Psithyrus males and females
disperse and mate. The males do not survive the winter but, like
nonparasitic bumblebee queens, Psithyrus females find suitable
locations to spend the winter and enter diapause after mating. They usually emerge from hibernation
later than their host species. Each species of cuckoo bee has a specific host species, which it may
physically resemble.[85] In the case of the parasitism of B. terrestris by B. (Psithyrus) vestalis, genetic
analysis of individuals captured in the wild showed that about 42% of the host species' nests at a single
location[a] had "[lost] their fight against their parasite".[79]
Sting
Queen and worker bumblebees can sting. Unlike in honeybees, a bumblebee's stinger lacks barbs, so the
bee can sting repeatedly without injuring itself; by the same token, the stinger is not left in the
wound.[86][87] Bumblebee species are not normally aggressive, but may sting in defence of their nest, or
if harmed. Female cuckoo bumblebees aggressively attack host colony members, and sting the host
queen, but ignore other animals unless disturbed.[88]
Predators, parasites, and pathogens

Bumblebees, despite their ability to sting, are eaten by certain
predators. Nests may be dug up by badgers and eaten whole,
including any adults present.[89] Adults are preyed upon by robber
flies and beewolves in North America.[90] In Europe, birds including
bee-eaters and shrikes capture adult bumblebees on the wing;
smaller birds such as great tits also occasionally learn to take
bumblebees, while camouflaged crab spiders catch them as they visit
flowers.[91]
Bumblebee nest dug up and The great grey shrike is able to detect flying bumblebees up to 100 m
destroyed by a predator, probably a (330 ft) away; once captured, the sting is removed by repeatedly
badger squeezing the insect with the mandibles and wiping the abdomen on
a branch.[92] The European honey buzzard follows flying bees back
to their nest, digs out the nest with its feet, and eats larvae, pupae
and adults as it finds them.[93]
Bumblebees are parasitised by tracheal mites, Locustacarus buchneri; protozoans

including Crithidia bombi and Apicystis bombi; and microsporidians including
Nosema bombi and Nosema ceranae. The tree bumblebee B. hypnorum has
spread into the United Kingdom despite hosting high levels of a nematode that
normally interferes with queen bees' attempts to establish colonies.[94] Deformed
wing virus has been found to affect 11% of bumblebees in Great Britain.[95]
Female bee moths (Aphomia sociella) prefer to lay their eggs in bumblebee nests.
The A. sociella larvae will then feed on the eggs, larvae, and pupae left
unprotected by the bumblebees, sometimes destroying large parts of the nest.[96]
Relationship to humans Bumblebee stored as

food by a great grey
shrike
Agricultural use
Bumblebees are important pollinators of both crops and

wildflowers.[97] Because bumblebees do not overwinter the entire
colony, they do not stockpile honey, and therefore are not useful as
honey producers. Bumblebees are increasingly cultured for
agricultural use as pollinators, among other reasons because they
can pollinate plants such as tomato in greenhouses by buzz
pollination whereas other pollinators cannot.[98] Commercial
production began in 1987, when Roland De Jonghe founded the
Biobest company; in 1988 they produced enough nests to pollinate
Bumblebees and human culture:
40 hectares of tomatoes. The industry grew quickly, starting with
Bombus anachoreta on a Russian
other companies in the Netherlands. Bumblebee nests, mainly of
postage stamp, 2005
buff-tailed bumblebees, are produced in at least 30 factories around
the world; over a million nests are grown annually in Europe; Turkey
is a major producer.[99]
Bumblebees are Northern Hemisphere animals. When red clover was introduced as a crop to New
Zealand in the nineteenth century, it was found to have no local pollinators, and clover seed had
accordingly to be imported each year. Four species of bumblebee from the United Kingdom were
therefore imported as pollinators. In 1885 and 1886 the Canterbury Acclimatization Society brought in
442 queens, of which 93 survived and quickly multiplied. As planned, red clover was soon being
produced from locally-grown seed.[36] Bumblebees are also reared commercially to pollinate tomatoes
grown in greenhouses.[52] The New Zealand population of buff-tailed bumblebees began colonising
Tasmania, 1,500 miles (2,400 km) away, after being introduced there in 1992 under unclear
circumstances.[100]
Some concerns exist about the impact of the international trade in mass-produced bumblebee colonies.
Evidence from Japan[101] and South America[102] indicates bumblebees can escape and naturalise in new
environments, causing damage to native pollinators. Greater use of native pollinators, such as Bombus
ignitus in China and Japan, has occurred as a result.[103] In addition, mounting evidence indicates mass-
produced bumblebees may also carry diseases, harmful to wild bumblebees[104][105] and honeybees.[105]
In Canada and Sweden it has been shown that growing a mosaic of different crops encourages
bumblebees and provides higher yields than does a monoculture of oilseed rape, despite the fact that the
bees were attracted to the crop.[106]
Population decline
Bumblebee species are declining in Europe, North America, and Asia due to a number of factors,
including land-use change that reduces their food plants. In North America, pathogens are possibly
having a stronger negative effect especially for the subgenus Bombus.[107] A major impact on bumblebees
was caused by the mechanisation of agriculture, accelerated by the urgent need to increase food
production during the Second World War. Small farms depended on horses to pull implements and carts.
The horses were fed on clover and hay, both of which were permanently grown on a typical farm. Little
artificial fertiliser was used. Farms thus provided flowering clover and flower-rich meadows, favouring
bumblebees. Mechanisation removed the need for horses and most of the clover; artificial fertilisers
encouraged the growth of taller grasses, outcompeting the meadow flowers. Most of the flowers, and the
bumblebees that fed on them, disappeared from Britain by the early 1980s. The last native British short-
haired bumblebee was captured near Dungeness in 1988.[108] This significant increase in pesticide and
fertilizer use associated with the industrialization of agriculture has had adverse effects on the genus
Bombus. The bees are directly exposed to the chemicals in two ways: by consuming nectar that has been
directly treated with pesticide, or through physical contact with treated plants and flowers. The species
Bombus hortorum in particular has been found to be impacted by the pesticides; their brood
development has been reduced and their memory has been negatively affected. Additionally, pesticide
use negatively impacts colony development and size.[109]
Bumblebees are in danger in many developed countries due to habitat destruction and collateral
pesticide damage. The European Food Safety Authority ruled that three neonicotinoid pesticides
(clothianidin, imidacloprid, and thiamethoxam) presented a high risk for bees.[110] While most work on
neonicotinoid toxicity has looked at honeybees, a study on B. terrestris showed that "field-realistic"
levels of imidacloprid significantly reduced growth rate and cut production of new queens by 85%,
implying a "considerable negative effect" on wild bumblebee populations throughout the developed
world.[111] However, in another study, following chronic exposure to field-realistic levels of the
neonicotinoid pesticide thiamethoxam, colony weight gain was not affected, nor were the number or
mass of sexuals produced.[112] Low levels of neonicotinoids can reduce the number of bumblebees in a
colony by as much as 55%, and cause dysfunction in the bumblebees' brains. The Bumblebee
Conservation Trust considers this evidence of reduced brain function "particularly alarming given that
bumblebees rely upon their intelligence to go about their daily tasks."[113] A study on B. terrestris had
results that suggests that use of neonicotinoid pesticides can affect how well bumblebees are able to
forage and pollinate. Bee colonies that had been affected by the pesticide released more foragers and
collected more pollen than bees who had not been dosed with neonicotinoid.[114] Although the bees
affected by the pesticide were able to collect more pollen, they took a longer amount of time doing so.[115]
Of 19 species of native nestmaking bumblebees and six species of cuckoo bumblebees formerly
widespread in Britain,[116] three have been extirpated,[117][118] eight are in serious decline, and only six
remain widespread.[119] Similar declines have been reported in Ireland, with four species designated
endangered, and another two considered vulnerable to extinction.[120] A decline in bumblebee numbers
could cause large-scale changes to the countryside, resulting from inadequate pollination of certain
plants.[121]
Some bumblebees native to North America are also vanishing, such as Bombus balteatus,[122] Bombus
terricola,[123] Bombus affinis,[124][125] and Bombus occidentalis, and one, Bombus franklini, may be
extinct.[126] In South America, Bombus bellicosus was extirpated in the northern limit of its distribution
range, probably due to intense land use and climate change effects.[127]
Conservation efforts
In 2006 the bumblebee researcher Dave Goulson founded a

registered charity, the Bumblebee Conservation Trust, to prevent the
extinction "of any of the UK's bumblebees."[128][129] In 2009 and
2010, the Trust attempted to reintroduce the short-haired
bumblebee, Bombus subterraneus, which had become extinct in
Britain, from the British-derived populations surviving in New
Zealand from their introduction there a century earlier.[130] From
2011 the Trust, in partnership with Natural England, Hymettus and
the RSPB, has reintroduced short-haired bumblebee queens from
Skåne in southern Sweden to restored flower-rich meadows at
Dungeness in Kent. The queens were checked for mites and
American foulbrood disease. Agri-environment schemes spread
Drone short-haired bumblebee, across the neighbouring area of Romney Marsh have been set up to
Bombus subterraneus. The species provide over 800 hectares of additional flower-rich habitat for the
was successfully reintroduced to bees. By the summer of 2013, workers of the species were found near
England from Sweden. the release zone, proving that nests had been established. The
restored habitat has produced a revival in at least five "Schedule 41
priority" species: the ruderal bumblebee, Bombus ruderatus; the
red-shanked carder bee, Bombus ruderarius; the shrill carder bee, Bombus sylvarum; the brown-
banded carder bee, Bombus humilis and the moss carder bee, Bombus muscorum.[131]
The world's first bumblebee sanctuary was established at Vane Farm in the Loch Leven National Nature
Reserve in Scotland in 2008.[121] In 2011, London's Natural History Museum led the establishment of an
International Union for the Conservation of Nature Bumblebee Specialist Group, chaired by Dr. Paul H.
Williams,[132] to assess the threat status of bumblebee species worldwide using Red List criteria.[133]
Bumblebee conservation is in its infancy in many parts of the world, but with the realization of the
important part they play in pollination of crops, efforts are being made to manage farmland better.
Enhancing the wild bee population can be done by the planting of wildflower strips, and in New Zealand,
bee nesting boxes have achieved some success, perhaps because there are few burrowing mammals to
provide potential nesting sites in that country.[106]
Misconception about flight
According to 20th-century folklore, the laws of aerodynamics prove

the bumblebee should be incapable of flight, as it does not have the
capacity (in terms of wing size or beats per second) to achieve flight
with the degree of wing loading necessary.[135]
'Supposedly someone did a back of the envelope

calculation, taking the weight of a bumblebee and its
wing area into account, and worked out that if it only
flies at a couple of metres per second, the wings wouldn't A widely believed falsehood holds
produce enough lift to hold the bee up,' explains Charlie that scientists proved bumblebees
Ellington, Professor of Animal Mechanics at Cambridge to be incapable of flight.[134]
University.[135]
The origin of this claim has been difficult to pin down with any certainty. John H. McMasters recounted
an anecdote about an unnamed Swiss aerodynamicist at a dinner party who performed some rough
calculations and concluded, presumably in jest, that according to the equations, bumblebees cannot
fly.[136] In later years, McMasters backed away from this origin, suggesting there could be multiple
sources, and the earliest he has found was a reference in the 1934 book Le Vol des Insectes by French
entomologist Antoine Magnan (1881–1938); they had applied the equations of air resistance to insects
and found their flight was impossible, but "One shouldn't be surprised that the results of the calculations
don't square with reality".[137]
The following passage appears in the introduction to Le Vol des Insectes:[138]
Tout d'abord poussé par ce qui se fait en aviation, j'ai appliqué aux insectes les lois de la
résistance de l'air, et je suis arrivé avec M. Sainte-Laguë à cette conclusion que leur vol est
impossible.
This translates to:
First prompted by what is done in aviation, I applied the laws of air resistance to insects,
and I arrived, with Mr. Sainte-Laguë, at this conclusion that their flight is impossible.
Magnan refers to his assistant André Sainte-Laguë.[139] Some credit physicist Ludwig Prandtl (1875–
1953) of the University of Göttingen in Germany with popularizing the idea. Others say Swiss gas
dynamicist Jakob Ackeret (1898–1981) did the calculations.[140]
The calculations that purported to show that bumblebees cannot fly

are based upon a simplified linear treatment of oscillating aerofoils.
The method assumes small amplitude oscillations without flow
separation. This ignores the effect of dynamic stall (an airflow
separation inducing a large vortex above the wing), which briefly
produces several times the lift of the aerofoil in regular flight. More
sophisticated aerodynamic analysis shows the bumblebee can fly
because its wings encounter dynamic stall in every oscillation
Bumblebee in flight. It has its tongue
cycle.[141]
extended and a laden pollen basket.
Additionally, John Maynard Smith, a noted biologist with a strong
background in aeronautics, has pointed out that bumblebees would
not be expected to sustain flight, as they would need to generate too
much power given their tiny wing area. However, in aerodynamics experiments with other insects, he
found that viscosity at the scale of small insects meant even their small wings can move a very large
volume of air relative to their size, and this reduces the power required to sustain flight by an order of
magnitude.[142]
In music and literature
The orchestral interlude Flight of the Bumblebee was composed (c. 1900) by Nikolai Rimsky-Korsakov. It
represents the turning of Prince Guidon into a bumblebee so he can fly away to visit his father, Tsar
Saltan, in the opera The Tale of Tsar Saltan,[143] although the music may reflect the flight of a bluebottle
rather than a bumblebee.[144] The music inspired Walt Disney to feature a bumblebee in his 1940
animated musical Fantasia and have it sound as if it were flying in all parts
of the theater. This early attempt at "surround sound" was unsuccessful,
and the music was excluded from the film's release.[145]
In 1599, during the reign of Queen Elizabeth I, someone, possibly Tailboys

Dymoke, published Caltha Poetarum: Or The Bumble Bee, under the
pseudonym "T. Cutwode".[146] This was one of nine books censored under
the Bishop's Ban issued by the Archbishop of Canterbury John Whitgift and
the Bishop of London Richard Bancroft.[147]
Emily Dickinson made a bumblebee the

subject of her parody of Isaac Watts's well-
known poem about honeybees, How Doth the
Little Busy Bee (1715). Where Watts wrote
The Russian composer "How skilfully she builds her cell! How neat
Nikolai Rimsky-Korsakov she spreads the wax!",[148] Dickinson's poem,
wrote the Flight of the
"The Bumble-Bee's Religion" (1881) begins
Bumblebee, c. 1900
"His little Hearse-like Figure / Unto itself a
Dirge / To a delusive Lilac / The vanity divulge
/ Of Industry and Morals / And every
righteous thing / For the divine Perdition / of Idleness and Spring." The
letter was said to have enclosed a dead bee.[149][150]
Emily Dickinson wrote a
The entomologist Otto Plath wrote Bumblebees and Their Ways in poem about a bumblebee.
1934.[151] His daughter, the poet Sylvia Plath, wrote a group of poems about Daguerreotype, c. 1848
bees late in 1962, within four months of her suicide,[152] transforming her
father's interest into her poetry.[153]
The scientist and illustrator Moses Harris (1731–1785) painted accurate

watercolour drawings of bumblebees in his An Exposition of English Insects
Including the Several Classes of Neuroptera, Hymenoptera, & Diptera, or
Bees, Flies, & Libellulae (1776–80).[154]
Bumblebees appear as characters, often eponymously, in children's books.

The surname Dumbledore in the Harry Potter series (1997–2007) is an old
name for bumblebee.[8] J. K. Rowling said the name "seemed to suit the
headmaster, because one of his passions is music and I imagined him
walking around humming to himself".[155] J. R. R. Tolkien, in his poem
Errantry, also used the name Dumbledor, but for a large bee-like creature.
Among the many books for younger children are Bumble the Bee by Yvon
Bumblebees of different Douran and Tony Neal (2014); Bertie Bumble Bee by K. I. Al-Ghani (2012);
species illustrated by Ben the Bumble Bee: How do bees make honey? by Romessa Awadalla
Moses Harris in his 1782(2015); Bumble Bee Bob Has a Big Butt by Papa Campbell (2012); Buzz,
Exposition of English
Buzz, Buzz! Went Bumble-bee by Colin West (1997); Bumble Bee by
Insects
Margaret Wise Brown (2000); How the Bumble Came to Bee by Paul and
Ella Quarry (2012); The Adventures of Professor Bumble and the Bumble
Bees by Stephen Brailovsky (2010). Among Beatrix Potter's "little books",
Babbity Bumble and other members of her nest appear in The Tale of Mrs. Tittlemouse (1910).
Military
The United States Naval Construction Battalions adopted the bumblebee as their insignia in 1942.
See also
Ophrys bombyliflora, the bumblebee orchid
Notes
a. The study location was the Botanical Garden Halle (Saale) in Germany, described as a flower-rich
region with high and stable abundance of both host and cuckoo species. 24 B. terrestris workers and
24 drones were captured on foraging flights. 24 male B. vestalis were similarly captured. DNA
analysis was used to estimate how many colonies these individuals came from.[79]
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scientists having proved that a bumblebee can't fly appears regularly in magazine and newspaper
stories. It's also the kind of item that can come up in a cocktail party conversation when the subject
turns to science or technology. [...] Often, the statement is made in a distinctly disparaging tone
aimed at putting down those know-it-all scientists and engineers who are so smart yet can't manage
to understand something that's apparent to everyone else. [...] the story has had remarkable staying
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refers to a calculation by his assistant André Sainte-Laguë, who was an engineer. The conclusion
was presumably based on the fact that the maximum possible lift produced by aircraft wings as small
as a bumblebee's wings and traveling as slowly as a bee in flight would be much less than the weight
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External links
Bumblebees of the world – find species by region, species groups, colour pattern (http://www.nhm.a
c.uk/research-curation/projects/bombus/), nhm.ac.uk
Bumblebee Conservation Trust (http://www.bumblebeeconservation.org/)
IUCN's Bumblebee Specialist Group (http://www.iucn.org/about/work/programmes/species/who_we_
are/ssc_specialist_groups_and_red_list_authorities_directory/invertebrates/bumblebee_specialist_gr
oup/)
Bombus Identification Guide (http://www.discoverlife.org/mp/20q?guide=Bombus), Discover Life: List
of Species (http://www.discoverlife.org/mp/20q?search=Bombus), Worldwide Species Map (http://ww
w.discoverlife.org/mp/20m?kind=Bombus_).
Deciphering the Mystery of Bee Flight (http://www.caltech.edu/news/deciphering-mystery-bee-flight-1

075)
Retrieved from "https://en.wikipedia.org/w/index.php?title=Bumblebee&oldid=971605560"
This page was last edited on 7 August 2020, at 04:30 (UTC).
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8/15/2020 Bumblebee - Wikipedia

Like their relatives the honeybees, bumblebees feed on nectar,

Etymology and common names

According to the Oxford English Dictionary (OED), the term "bumblebee"

An old provincial name, "dumbledor", also denoted a buzzing insect such as

Subgenera of the genus Bombus

Distribution and habitat

Many species of Bombus, including the group sometimes called Psithyrus

Cuckoo bumblebees, like

Communication and social learning

Reproduction and nesting

Nest size depends on species of bumblebee. Most form colonies of

After the emergence of the

Bumblebees generally visit flowers that exhibit the bee pollination

Bumblebees use a combination of colour and spatial relationships to

Pollen is removed from flowers deliberately or incidentally by

In at least some species, once a bumblebee has visited a flower, it

Asynchronous ﬂight muscles

Psithyrus female, such as that of the Psithyrus species of B.

The female Psithyrus has a number of morphological adaptations for

Predators, parasites, and pathogens

Bumblebees are parasitised by tracheal mites, Locustacarus buchneri; protozoans

Relationship to humans Bumblebee stored as

Bumblebees are important pollinators of both crops and

In 2006 the bumblebee researcher Dave Goulson founded a

Misconception about ﬂight

According to 20th-century folklore, the laws of aerodynamics prove

'Supposedly someone did a back of the envelope

The following passage appears in the introduction to Le Vol des Insectes:[138]

This translates to:

The calculations that purported to show that bumblebees cannot fly

In music and literature

In 1599, during the reign of Queen Elizabeth I, someone, possibly Tailboys

Emily Dickinson made a bumblebee the

The scientist and illustrator Moses Harris (1731–1785) painted accurate

Bumblebees appear as characters, often eponymously, in children's books.

85. Macdonald, 2003. p. 12

16. Kaae, Richard (nd). "Bees, Wasps and Ants" (https://web.archive.org/web/20131224121927/http://drk

33. "2011 Update" (https://web.archive.org/web/20121203034020/http://cmsdata.iucn.org/downloads/bu

Deciphering the Mystery of Bee Flight (http://www.caltech.edu/news/deciphering-mystery-bee-ﬂight-1

Retrieved from "https://en.wikipedia.org/w/index.php?title=Bumblebee&oldid=971605560"

This page was last edited on 7 August 2020, at 04:30 (UTC).

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